Response of invertebrates to lotic disturbance: Is the hyporheic zone a patchy refugium?

1. Natural experiments, in the form of disturbance from spates, were used to study the resistance and resilience of interstitial communities. Investigations were conducted in a by-passed section of the Rhone River characterized by an artificial hydrology with frequent spates separated by regular minimum discharge of 30 m(3) s(-1). 2. Three areas of a bar were studied, upwellings at the head of the bar (stations 1 and 2), and downwelling at the tail of the bar (station 3). In the head of the bar the substratum was characterized by stable cobbles, while mobile gravels dominated in the tail of the bar. At each station, samples were derived from four depths (0.5, 1.0, 1.5 and 2.0 m below the surface of the substratum). Fifteen spates occurred during the study period whose peak discharge ranged from 50 to 1640 m(3) s(-1). Temporal variations of the fauna were studied by comparing the spate effect observed 1 day (resistance), 7 days (resilience) and 17 days after the spate. Within-class correspondence analysis was used to compare the temporal variability of the fauna within each class (station/depth). 3. The fauna differed markedly between the three stations, and the relative density of stygobionts (i.e. hypogean fauna) decreased from 55% at station 1 to 4% at station 3. The spatio-temporal variability increased dramatically from station 1 to station 3. 4. The results suggest that the hyporheic zone acts as a patchy refugium: the stations were more or less active refugial zones, depending on hydrology (upwelling or downwelling), substratum stability and spate amplitude. 5. The downwelling station was the main refugium area for benthic taxa. Important migrations of benthic groups (e.g. Gammarus, Cladocera) or hyporheic taxa (e.g. Cyclopoida and Harpacticoida) were observed deep into the sediment (2 m). Vertical movements of stygobionts (Niphargus, Niphargopsis) were also observed at high amplitude spates. These movements were very important (great numbers of individuals migrated) at low and medium magnitude spates, but were unimportant at high discharge, when the threshold of sediment instability was exceeded. In this case the substratum became mobile and induced drift of benthic organisms. 6. Conversely, in the upwelling stable stations, accumulation was less important (lower number of species and lower densities) but more constant with increasing discharge, suggesting that substratum stability is also a key factor. 7. Generally recovery was rapid at all stations (within 7 days) but no relationships were found between resilience (rate of recovery) and the amplitude of spates.